Milenge Kamalebo and De Kesel Journal of Ethnobiology and Ethnomedicine (2020) 16:8 https://doi.org/10.1186/s13002-020-0357-5

RESEARCH Open Access Wild edible ectomycorrhizal fungi: an underutilized food resource from the rainforests of Tshopo province (Democratic Republic of the Congo) Héritier Milenge Kamalebo1,2* and André De Kesel3

Abstract Background: Ectomycorrhizal (EcM) fungi constitute a source of income as well as proper food with considerable nutritional value. Although edible EcM fungi are highly diverse and expected to host considerable nutritional attributes, only few studies focus on their use and promotion in the province of Tshopo (DR Congo). This study provides original ethnomycological and diversity data on edible ectomycorrhizal rainforest fungi from the Man-and- Biosphere reserve of Yangambi and the reserve of Yoko. Methods: The list of edible fungi follows the current . Taxa were collected in plots situated in different types of rainforests. Each taxon is supported by herbarium reference specimens. Ethnomycological data on locally consumed EcM fungi were collected from randomly selected people living near the Man-and-Biosphere reserve of Yangambi and the Yoko reserve. People were interviewed using a semi-structured questionnaire. The interview campaign involved 160 informants, all randomly selected from 6 different ethnic communities. Results: The results reveal that rainforests from the Yangambi Biosphere reserve and Yoko forest reserve provide a relatively high number of edible fungi, more than local people actually use. Mixed forest stands hold the highest diversity in saprotrophic edible fungi (p value < 0.001) while no significant difference (p value > 0.05) was observed in the number of saprotrophic and EcM fungi within monodominant forests. In spite of being accessible, this renewable natural resource is underexploited. Although a wide array of EcM fungi is available in primary forests dominated by ectomycorrhizal trees, local people’s major interest goes to the saprotrophic fungi from areas with degraded mixed forests. Conclusion: The lack of local interest for EcM fungi is probably related to the considerable distance people have to cover to collect them. As a result, the edible EcM fungi from the Tshopo area represent a potentially interesting but underutilized resource. Keywords: Edible ectomycorrhizal fungi, Fungal consumption, Tshopo, Democratic Republic of the Congo

* Correspondence: [email protected] 1Faculté des Sciences, Université de Kisangani, BP 2012, Kisangani, Democratic Republic of the Congo 2Centre de Recherches Universitaires du Kivu (CERUKI-ISP), BP 854, Bukavu, Democratic Republic of the Congo Full list of author information is available at the end of the article

© The Author(s). 2020 Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The Creative Commons Public Domain Dedication waiver (http://creativecommons.org/publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated. Milenge Kamalebo and De Kesel Journal of Ethnobiology and Ethnomedicine (2020) 16:8 Page 2 of 13

Introduction Tshopo is mainly characterized by a tropical evergreen In many regions of the world, including tropical Africa, rainforest and some groves of semi-deciduous forests edible fungi often constitute a source of income and food, found on hills and plateaus [20–22]. These rainforests with a non-negligible nutritional value [1–6]. According are mainly dominated by species such as Gilbertioden- to De Kesel et al. [4] and Degreef et al. [7], documenting dron dewevrei (De Wild.) J. Léonard, Brachystegia laur- the relative importance and potential of locally used EcM entii (De Wild.) Louis, Scorodophloeus zenkeri Harms, fungi remains vital for improving strategies for their con- Prioria balsamifera (Vermoesen) Breteler, and Julbernar- servation and promoting sustainable use. Edible EcM dia seretii (De Wild.) Troupin [20–24]. fungi provide high-quality and important amount of crude The fungi were collected between 2013 and 2016, protein, minerals and carbohydrates, fats, etc. [2, 3, 5, 6]. mostly within plots situated in the rainforests of the Bio- The amount of fungal crude protein for instance is gener- sphere reserve of Yangambi (0° 51′ 01.62″ N; 24° 31′ ally ranked between 19 and 35% of the dry mass [2]. Fur- 43.53″ E) and Yoko reserve (0° 17′ 34.9″ N; 25° 18′ thermore, many species of edible EcM fungi provide 27.4″ E) (Fig. 1). Monitoring of plots follows Lodge et al. higher amount of unsaturated fatty acids than saturated [25], and fungal sampling was performed within forests ones [2, 5]. Unlike saturated fatty acids (found in high dominated by Gilbertiodendron dewevrei, Brachystegia amount in animal fats), unsaturated fatty acids are sug- laurentii, Julbernardia seretii, Uapaca heudelotii, and gested to be essential in the human diet [2]. The high pro- Uapaca guineensis and in mixed forests. Three plots of portion of unsaturated fatty acids and high percentage of 100 × 100 m were demarcated in each type of forest; linoleic acid found in edible EcM mushrooms lead to con- each of them divided in a 20 × 20 m grid. Due to the sider them as a healthy food [2]. In addition, the proteins elongated shape of Uapaca heudelotii-dominated forests, of most of edible EcM taxa contain all nine essential more stretched plots were installed in this forest type. In amino acids (leucine, isoleucine, valine, tryptophan, lysine, each plot, the aboveground fruiting bodies of all EcM threonine, phenylalanine, methionine, and histidine) used fungi were collected by walking parallel bands covering in humans’ nutrition [2, 5]. the entire plot [26]. Unidentifiable fruit bodies were EcM fungi constitute one of the most important photographed in situ and dried after notes were taken groups of edible fungi worldwide. In tropical Africa, from their macromorphological features (following [18]). EcM fungi are massively consumed in the Zambezian re- Voucher specimens were dried using a field drier [27] gion [4, 5, 8–14], and in the savanna woodlands and and deposited at the herbarium of Meise Botanic Garden open forest areas of the Soudanian region [15–17]. (Belgium). Within the Zambezian geographical zone, local people collect, sell, and consume impressive quantities of EcM Fungal identification taxa. The socio-economic interest for EcM fungi is con- The identification of voucher specimens was done using sidered a major incentive for obtaining strong and macroscopic and microscopic characteristics, as outlined in broadly supported forest conservation programs [4]. Eyi-Ndong et al. [18]. The available taxonomic literature only Within the rainforests of Tshopo province (Demo- covers a fraction of the Central African rainforest fungi. The cratic Republic of the Congo), many species of edible following contributions were used for identification: Heine- EcM fungi are available. In this area, EcM fungi are typ- mann [28], Heim [29], Pegler [30], Heinemann and Ramme- ically found within primary forests dominated by EcM loo [31–33], Buyck [8, 34, 35],DeKeseletal.[4, 9, 16], partner trees such as Gilbertiodendron dewevrei (De Verbeken and Walleyn [36], Eyi-Ndong et al. [18], as well as Wild.) J. Léonard, Brachystegia laurentii (De Wild.) identification keys provided by the Flora of Tropical Louis, Julbernardia seretii (De Wild.) Troupin, Uapaca Africa (https://www.ffta-online.org/) and Edible Fungi of guineensis Müll. Arg, and U. heudelotii Baillon [18, 19]. Tropical Africa (https://www.efta-online.org/). Species names This study aims to assess the distribution and diversity and author’s abbreviations largely follow Index Fungorum of edible fungi within various types of rainforests found (http://www.indexfungorum.org/Names/Names.asp). Uniden- in the Man-and-Biosphere reserve of Yangambi and the tified taxa were left out of the analysis. Yoko reserve. We show how access and availability, as well as trophic level of fungi, affect the interest of local Ethnomycological data acquisition and treatment people, and how this in turn creates or annihilates op- Ethnomycological data on locally consumed fungi were col- portunities for forest conservation. lected from local people living around the Man-and- Biosphere reserve of Yangambi and the Yoko reserve. Data Materials and methods were collected using open, semi-structured interviews (paper Study site and sampling of fungi fill-in questionnaires). The interviews involved mainly the Fieldwork was done in the province of Tshopo, located head of the family, sometimes assisted by other family mem- 2° N 2° S and 22° E 28° E [20]. The vegetation from bers. The questions focused basically on the informant’s Milenge Kamalebo and De Kesel Journal of Ethnobiology and Ethnomedicine (2020) 16:8 Page 3 of 13

Fig. 1 Location of the study area

knowledge concerning the different locally consumed edible 600 ≈ 1þðNÂ0:072Þ 152, households that were fitted to 160 infor- fungi. Interviews were obtained from 160 informants, all ran- mants. Referring to the mean distribution of households per domly selected, but belonging to one of 6 ethnic communi- sampling site (villages and suburbs of Yangambi), 7 house- ties (Bakumu, Turumbu, Topoke, Lokele, Ngelema, and holds were interviewed from each of the 4 villages while 38 Ngando). The entire pool of informants counted 88 men fromEkuchu,30fromLusambila,20fromManzikala,and (55%) and 72 women (45%), ranging from 16 to 72 years old. 12 households from IFA. In each sampling site, all house- The interviewed communities live in four villages in the holds were numbered. The first numbers referring to the vicinity of the Man-and-Biosphere reserve of Yangambi considered sample size were selected randomly using the (Yakako, Yalungu, Lyoli, and Lobiloto), 4 suburbs of Yan- function “rand.between” of the Excel software. gambi city (IFA, Lusambila, Ekuchu, and Manzikala), and 4 The analyzed data only refer to locally eaten fungi and villages surrounding the Yoko forest reserve (Babogombe, allow to present (per species) information on the edible Biaro, PK 48, and PK 25). mushrooms’ cultural significance (EMCS). According to With an average of 600 households living in the studied Pieroni [38], the edible mushrooms’ cultural significance area, the average number of studied households revolves index refers to the importance or the role that a given fungal around 25 per village (4.2% of the entire pool). Within the 4 taxon or group of fungi plays in the social life of a group of suburbs of Yangambi, the distribution of households was 150 people or a community. Using pictures or fresh sporocarps (25%), 120 (20%), 80 (13.3%), and 50 (8.3%) respectively re- of edible fungi, the edible mushrooms’ cultural significance ported from Ekuchu, Lusambila, Manzikala, and IFA. Ac- for a fungal species corresponds with the sum of the scores cording to Gumucio et al. [37],thesamplesize(n)ofthe for “edibility status” given by all informants, divided by the interviewed households should be calculated as follows: n = total number of informants. Edibility status scores or fre- N 1þNÂe2 ,whereN is the total number of available households quencies of mention (FM) were assigned through informants’ and e is the level of precision. With a precision level of ± 7%, answers to the following question: Do you eat this mush- thesamplesizewascalculatedbythefollowingformula:n = room? (yes = 1, no = 0). A Kruskal-Wallis analysis was used Milenge Kamalebo and De Kesel Journal of Ethnobiology and Ethnomedicine (2020) 16:8 Page 4 of 13

to test the role of ethnicity on the edibility status score, and eaten by informants from different ethnic groups how this changes according to the different trophic groups. (Table 3). Because of their low numbers (three species), differences in the mean numbers of consumed termite- Results associated species are evidently small. In general, the Checklist and diversity of wild edible fungi most commonly eaten mushrooms (with the highest Table 1 presents the checklist of edible fungi recorded edibility index) are saprotrophic or woody-decaying from the rainforests of Tshopo. Based on the informa- fungi (Table 3), the most appreciated ones being Auri- tion found in the literature and ethnomycological inquir- cularia spp., Marasmius buzungolo, Lentinus squarro- ies, 78 species of macrofungi are edible. sulus,andSchizophyllum commune (Fig. 4). Only a small fraction of EcM fungi, especially from the genus Trophic groups and distribution of edible fungi Cantharellus, are reported as a delicacy by some popu- Edible species belong to 3 trophic groups (saprotrophic, lations. The scores of EcM fungi are even more low- ectomycorrhizal, and termite associated). The sapro- ered, knowing that some local populations (Kumu) trophic taxa (wood and litter decaying fungi) represent systematically reject all edible Russula because of their the most species-rich group (44 species), followed by the vivid colors. In terms of diversity and quantity, local EcM fungi (31 species) and termite-associated taxa (3 markets (Yangambi) offer far more saprotrophic than species, all Termitomyces). Whereas the number of ed- EcM fungi (De Kesel pers. obs.). ible fungi significantly differs between trophic groups, strong variation in the composition of edible fungi is ob- Discussion served between forest types (Fig. 2). Figure 2 shows that This study revealed that wild edible fungi from rainfor- the mixed forests, on average, hold the highest diversity ests of Tshopo belong to several trophic groups. They in saprotrophic edible fungi, while EcM fungi reach their can be either saprotrophic (growing on dead fallen highest species numbers in the monodominant forests. trunks and litter), or termite associated (growing in mu- Edible termite-associated fungi (Termitomyces sp.) show tualistic relation with termites), or ectomycorrhizal, i.e., the lowest species numbers and seem equally repre- living in a mutualistic relation with the roots of some sented in both mixed and monodominant forests. Con- vascular plants. Within rainforests from the Yangambi sidering all fungal trophic groups, plots from Biosphere reserve and Yoko reserve, relatively high num- monodominant and mixed forests showed similar aver- bers of edible fungi are available. A more quantitative age numbers of edible species, but variations are more approach, similar to the one used by Yorou et al. [41]or important in the monodominant forests. The result is De Kesel et al. [4], is needed to ascertain this, but it that the highest numbers of edible species were encoun- seems that the EcM fraction of edible fungi is still un- tered in some of the monodominant forests, especially in derexploited. Although a wide array of edible EcM fungi the forest dominated by Gilbertiodendron dewevrei. is available in the primary ectomycorrhizal forests, the Within monodominant forests, we observed no signifi- interest of local people mainly goes to saprotrophic fungi cant difference in the number of saprotrophic and EcM from the degraded mixed forests. Milenge et al. [39] fungi (Fig. 3). In contrast, within mixed forests, the num- reported that most of these saprotrophic edible taxa typ- ber of saprotrophic taxa was significantly higher than in ically grow on wood (logs). the EcM forests. Among the reported edible EcM fungi, only species of the genus Cantharellus are considered of some im- Usage of edible fungi portance in local consumption [39]. As stated by Eyi- Almost a third (25 species) of the 78 edible species are not Ndong et al. [18], species of the genus Cantharellus used for food by any of the interviewed people (Table 2). are the most important edible EcM fungi found This is the case for Cantharellus conspicuus, C. incarnatus, within rainforests from the central African Congo Cookeina tricholoma, Coprinus plicatilis, Dacryopinax basin. Most species of Cantharellus are known world- spathularia, Gerronema hungo, Hygrocybe coccinea, Hypho- wide as good or even excellent edibles. With its 42 loma subviride, Lactifluus gymnocarpus, L. heimii, L. African endemic species, the genus is by far best rep- pelliculatus, Lepista sordida, Leucocoprinus discoideus, resented on the African continent [9]. In contrast to Macrolepiota dolichaula, Marasmiellus inoderma, Neonotho- the mushroom diet chosen by people from Tshopo, panus hygrophanus, Paxillus brunneotomentosus, Polyporus Cantharellus species are massively consumed in the arcularius, Psathyrella candolleana, Russula roseostriata, R. Zambezian region [4, 5, 9, 10, 13, 14, 42]. sese, Tremella fuciformis, Tylopilus balloui, Volvariella par- Although other EcM genera, i.e., Amanita, , vispora,andVolvopluteus gloiocephalus. Lactifluus,andRussula, are common in the forests domi- Significant differences were observed in the mean nated by Gilbertiodendron dewevrei, these edible EcM number of ectomycorrhizal and saprotrophic fungi fungi do not seem to be at all harvested and consumed by Milenge Kamalebo and De Kesel Journal of Ethnobiology and Ethnomedicine (2020) 16:8 Page 5 of 13

Table 1 List of edible fungi from rainforests of Tshopo Species Edibility Trophic FM EMCS Supporting literature Voucher status group score specimens 1. Agaricus bambusicola Heinem. Food Sapr 24 0.2 Milenge et al. [39] ADK5267 2. Agaricus crocopeplus Berk. & Broome Food Sapr 27 0.2 Milenge et al. [39] ADK5747, MKH073 3. Amanita annulatovaginata Beeli Food EcM 41 0.3 Milenge et al. [39] MKH016 4. Amanita echinulata Beeli Food EcM 35 0.2 Milenge et al. [39] ADK5938, MKH159 5. Amanita pudica (Beeli) Walleyn Food EcM 49 0.3 Degreef et al. [7], De Kesel et al. [4], ADK5924 Milenge et al. [39] 6. Amanita robusta Beeli Food EcM 49 0.3 Boa [1], Milenge et al. [39] ADK5236 7. Armillaria heimii Pegler Food Sapr 78 0.5 Eyi-Ndong et al. [18], Degreef et al. [7], ADK5230, Milenge et al. [39] MKH046 8. Auricularia cornea Ehrenb. Food Sapr 144 0.9 De Kesel et al. [16], Boa [1], Eyi-Ndong et al. [18], ADK5175 Degreef et al. [7], De Kesel et al. [4], Milenge et al. [39] 9. Auricularia delicata (Mont.ex Fr.) Henn. Food Sapr 142 0.9 De Kesel et al. [16], Boa [1], Degreef et al. [7], ADK5169, De Kesel et al. [4], Milenge et al. [39] MKH214 10. Calyptella longipes (Cooke & Massee) W.B. Food Sapr 12 0.1 Milenge et al. [39] ADK5868 Cooke 11. Cantharellus congolensis Beeli Food EcM 89 0.6 De Kesel et al. [16], Boa [1], Eyi-Ndong et al. [18], ADK5199, Degreef et al. [7], De Kesel et al. [4], Milenge et al. [39] MKH180 12. Cantharellus conspicuus Eyssartier, Buyck & Edible EcM 0 0.0 Eyssartier et al. [40] ADK5937 Verbeken 13. Cantharellus densifolius Heinem. Food EcM 53 0.3 Boa [1], Eyi-Ndong et al. [18], Degreef et al. [7], MKH017 De Kesel et al. [4], Milenge et al. [39] 14. Cantharellus floridulus Heinem. Food EcM 54 0.3 De Kesel et al. [16], Boa [1], Eyi-Ndong et al. [18], ADK5670 De Kesel et al. [4], Milenge et al. [39] 15. Cantharellus incarnatus (Beeli) Heinem. Edible EcM 0 0.0 Boa [1], Eyi-Ndong et al. [18] MKH069, MKH166 16. Cantharellus longisporus Heinem. Food EcM 132 0.8 Boa [1], Milenge et al. [39] ADK5684, MKH141 17. Cantharellus luteopunctatus (Beeli) Heinem. Food EcM 51 0.3 Boa [1], Eyi-Ndong et al. [18], Milenge et al. [39] ADK5836, MKH078 18. Cantharellus miniatescens Heinem. Food EcM 39 0.2 [1], Eyi-Ndong et al. [18], Milenge et al. [39] ADK5216, MKH142 19. Cantharellus aff. ruber Heinem. Food EcM 29 0.2 Boa [1], Degreef et al. [7], De Kesel et al. [4], ADK5749 Milenge et al. [39] 20. Cantharellus rufopunctatus (Beeli) Heinem. Food EcM 84 0.5 Boa [1], Eyi-Ndong et al. [18], De Kesel et al. [4], ADK5892, Milenge et al. [39] MKH051 21. Cookeina speciosa (Fr.) Dennis Food Sapr 11 0.1 Eyi-Ndong et al. [18], Milenge et al. [39] MKH054 22. Cookeina tricholoma (Mont.) Kuntze Edible Sapr 0 0.0 Boa [1] ADK5701 23. Coprinellus disseminatus (Pers.) J.E. Lange Food Sapr 18 0.1 Milenge et al. [39] ADK5615 24. Coprinus plicatilis (Curtis) Fr. Edible Sapr 0 0.0 [1] – 25. Cotylidia aurantiaca (Pat.) A. L. Welden Food Sapr 14 0.1 Boa [1], Degreef et al. [7], Milenge et al. [39] ADK5625, MKH058 26. Dacryopinax spathularia (Schwein.) G.W. Edible Sapr 0 0.0 Boa [1], De Kesel et al. [4] ADK5578 Martin 27. Favolus tenuiculus (P. Beauv.) Fr. Food Sapr 8 0.1 Eyi-Ndong et al. [18], De Kesel et al. [4], MKH109, Milenge et al. [39] MKH195 28. Gerronema hungo (Henn.) Degreef & Eyi Edible Sapr 0 0.0 Eyi-Ndong et al. [18] MKH057, MKH201 29. Gymnopilus zenkeri (Henn.) Singer Food Sapr 83 0.5 Eyi-Ndong et al. [18], De Kesel et al. [4], ADK5619 Milenge et al. [39] Milenge Kamalebo and De Kesel Journal of Ethnobiology and Ethnomedicine (2020) 16:8 Page 6 of 13

Table 1 List of edible fungi from rainforests of Tshopo (Continued) Species Edibility Trophic FM EMCS Supporting literature Voucher status group score specimens 30. Hygrocybe cantharellus (Schwein.) Food Sapr 28 0.2 Boa [1], Milenge et al. [39] ADK5849, Murrill MKH229 31. Hygrocybe coccinea (Schaeff.) P. Edible Sapr 0 0.0 Boa [1] ADK5864 Kummer 32. Hymenagaricus luteolosporus Food Sapr 18 0.1 Milenge et al. [39] ADK5222 Heinem.& Little Flower 33. Hypholoma subviride Berk. Edible Sapr 0 0.0 Degreef et al. [7] ADK5891 & M.A. Curtis 34. Lactarius acutus Heim Food EcM 49 0.3 Eyi-Ndong et al. [18], Milenge et al. [39] ADK5251, MKH001 35. Lactifluus annulatoangustifolius Food EcM 28 0.2 Boa [1], Milenge et al. [39] ADK5862 (Beeli) Buyck 36. Lactifluus gymnocarpus Edible EcM 0 0.0 Boa [1], Eyi-Ndong et al. [18], De Kesel et al. [4] MKH135 (R. Heim ex Singer) Verbeken 37. Lactifluus heimii (Verbeken) Verbeken Edible EcM 0 0.0 Boa [1], De Kesel et al. [4] MKH170 38. Lactifluus pelliculatus (Beeli) Buyck Edible EcM 0 0.0 Eyi-Ndong et al. [18], Boa [1] MKH012 39. Lentinus squarrosulus Mont. Food Sapr 152 1.0 De Kesel et al. [16], Boa [1], Eyi-Ndong et al. [18], ADK5226, De Kesel et al. [4], Milenge et al. [39] MKH079 40. Lepista rhodotoides Singer Food Sapr 54 0.3 Milenge et al. [39] MKH085 41. Lepista sordida (Schumach.) Singer Edible Sapr 0 0.0 Boa [1], Degreef et al. [7], De Kesel et al. [4] ADK5254 42. Leucocoprinus cepaestipes (Sowerby) Pat Food Sapr 19 0.1 Boa [1], Milenge et al. [39] ADK5206 43. Leucocoprinus discoideus (Beeli) Heinem Edible Sapr 0 0.0 Boa [1] ADK5606 44. Macrolepiota dolichaula (Berk. & Broome) Edible Sapr 0 0.0 Boa [1], Eyi-Ndong et al. [18], Degreef et al. [7], ADK5207, Pegler & R.W. Rayner De Kesel et al. [4] MKH223 45. Marasmiellus inoderma (Berk.) Singer Edible Sapr 0 0.0 Eyi-Ndong et al. [18], Degreef et al. [7] ADK5737, MKH094 46. Marasmius arborescens (Henn.) Beeli Food Sapr 99 0.6 Boa [1], Eyi-Ndong et al. [18], [7], De Kesel et al. [4], ADK5255, Milenge et al. [39] MKH231 47. Marasmius bekolacongoli Beeli Food Sapr 62 0.4 Eyi-Ndong et al. [18], Degreef et al. [7], ADK5779, De Kesel et al. [4], Milenge et al. [39] MKH035 48. Marasmius buzungolo Singer Food Sapr 138 0.9 Boa [1], Eyi-Ndong et al. [18], Milenge et al. [39] ADK5760, MKH121 49. Marasmius confertus Berk. & Broome Food Sapr 51 0.3 Milenge et al. [39] ADK5276, MKH021 50. Neonothopanus hygrophanus (Mont.) De Edible Sapr 0 0.0 Eyi-Ndong et al. [18], De Kesel et al. [4] ADK5260, Kesel & Degreef MKH031 51. Paxillus brunneotomentosus Heinem. & Edible Sapr 0 0.0 Degreef et al. [7] ADK5655, Rammeloo MKH129 52. Pleurotus cystidiosus O.K. Mill. Food Sapr 118 0.7 De Kesel et al. [16], Boa [1], Degreef et al. [7], MKH067 Milenge et al. [39] 53. Pleurotus flabellatus (Berk. & Br.) Sacc. Food Sapr 48 0.3 Eyi-Ndong et al. [18], Degreef et al. [7], ADK5771 Milenge et al. [39] 54. Pleurotus tuber-regium (Fr.) Fr. Food Sapr 98 0.6 De Kesel et al. [16], Boa [1], Eyi-Ndong et al. [18], MKH106 Degreef et al. [7], De Kesel et al. [4], Milenge et al. [39] 55. Polyporus arcularius (Batsch) Fr. Edible Sapr 0 0.0 Boa [1] MKH107 56. Psathyrella candolleana (Fr.) Maire Edible Sapr 0 0.0 Boa [1] ADK5252 57. Rubinoboletus luteopurpureus (Beeli) Food EcM 24 0.2 Boa [1], Milenge et al. [39] ADK5192, Heinem. & Rammeloo MKH235 58. Russula annulata R. Heim Food EcM 32 0.2 Milenge et al. [39] MKH014 59. Russula inflata Buyck Food EcM 29 0.2 Milenge et al. [39] ADK5217 60. Russula meleagris Buyck Food EcM 35 0.2 Boa [1], Milenge et al. [39] MKH113 Milenge Kamalebo and De Kesel Journal of Ethnobiology and Ethnomedicine (2020) 16:8 Page 7 of 13

Table 1 List of edible fungi from rainforests of Tshopo (Continued) Species Edibility Trophic FM EMCS Supporting literature Voucher status group score specimens 61. Russula porphyrocephala Buyck Food EcM 26 0.2 Milenge et al. [39] ADK5750, MKH178 62. Russula pruinata Buyck Food EcM 27 0.2 Milenge et al. [39] MKH115, MKH151 63. Russula roseostriata Buyck Edible EcM 0 0.0 Boa [1], Eyi-Ndong et al. [18] MKH150 64. Russula sese Beeli Edible EcM 0 0.0 Boa [1], Eyi-Ndong et al. [18] MKH114 65. Russula sesemoindu Beeli Food EcM 49 0.3 Eyi-Ndong et al. [18], Milenge et al. [39] MKH146 66. Russula striatoviridis Buyck Food EcM 52 0.3 Boa [1], Eyi-Ndong et al. [18], Milenge et al. [39] ADK5896 67. Schizophyllum commune Fr. Food Sapr 160 1.0 De Kesel et al. [16], Boa [1], Eyi-Ndong et al. [18], MKH108 Degreef et al. [7], De Kesel et al. [4], Milenge et al. [39] 68. Tapinella panuoides (Fr.) E.-J. Gilbert Food Sapr 26 0.2 Milenge et al. [39] MKH043 69. Termitomyces robustus (Beeli) Heim Food Term 128 0.8 De Kesel et al. [16], Boa [1], Eyi-Ndong et al. [18], ADK5242 Degreef et al. [7], De Kesel et al. [4], Milenge et al. [39] 70. Termitomyces singidensis Saarim. & Härk. Food Term 66 0.4 Boa [1] ADK5852 71. Termitomyces striatus (Beeli) Heim Food Term 124 0.8 De Kesel et al. [16], Boa [1], Eyi-Ndong et al. [18], ADK5179 Degreef et al. [7], De Kesel et al. [4] 72. Tremella fuciformis Berkeley Edible Sapr 0 0.0 Boa [1] MKH047 73. Tricholomopsis aurea (Beeli) Desjardin & B.A. Food Sapr 64 0.4 Boa [1], Eyi-Ndong et al. [18] ADK5625, Perry. MKH154 74. Trogia infundibuliformis Berk. & Br. Food Sapr 38 0.2 Eyi-Ndong et al. [18], De Kesel et al. [4], ADK5223 Milenge et al. [39] 75. Tylopilus balloui (Peck) Singer Edible EcM 0 0.0 Boa [1] ADK5950 76. Volvariella parvispora Heinem. Edible Sapr 0 0.0 Boa [1], De Kesel et al. [4] ADK5786 77. Volvopluteus gloiocephalus (DC) Vizzini Edible Sapr 0 0.0 Boa [1] ADK5641 78. Xerocomus spinulosus Heinem. & Gooss.- Food EcM 12 0.1 De Kesel et al. [4], Milenge et al. [39] ADK5209 Font. Edible = species known as edible but not reported locally consumed, food = species locally consumed, FM frequency of mention

local populations of the Tshopo province. However, in from these NTFP goes hand in hand with sustainable miombo areas of Burundi [7, 8, 43], and similar woodlands use and preservation of host trees and their immediate in the Zambezian [4, 11] and Guineo-Soudanian ecozone environment. Losing revenue from not being able to [15–17], these genera (and Cantharellus)representa collect EcM fungi is a strong incentive to preserve, or major source of food and income. at least make sure, that the forest ecosystem keeps on Triggering the interest of local people for non- delivering its services. timber forest products requires science-based infor- In this context, it surely is interesting to monitor nat- mation about how much of the resource is available, ural productions of edible EcM fungi in different types how much and where it can be obtained in a sustain- of rainforests of the Democratic Republic of the Congo. able way, and what its value may be. Based on prod- So far, this has not been done. Gathering such data is uctivity data of the 50 most common edible species, one thing, but convincing local people of changing or 1 ha of Katangese miombo woodland produces an widening their diet, i.e., to include more EcM fungi, is average of 148 kg of fresh fruiting bodies per year [4]. another. Several scientists [7, 17, 27, 44–47] have re- A very similar biomass, i.e., 140.7 kg/ha/year, was ob- portedtheroleofcultureandtraditiononthevariety tained from 29 edible species monitored in the sa- of mushrooms consumed by local people. In the vanna woodlands of Benin [17]. From miombo context of the province of Tshopo, taboos resting on woodlands of Katanga in the Democratic Republic of mushroom consumption create mycophobia and subse- the Congo for instance, 1 kg of fresh fruiting bodies of quently decrease the interest for some edible EcM spe- Cantharellus spp. costs approximatively 2.8 USD [4]. cies. In spite of their high nutritional value, most Local people realize that the yearly revenue obtained Russula species are not eaten by Kumu people because Milenge Kamalebo and De Kesel Journal of Ethnobiology and Ethnomedicine (2020) 16:8 Page 8 of 13

Fig. 2 Diversity of edible fungi according to trophic groups and forest stands

of vivid colors of the cap. Variation in mushroom con- sumption may also rely on the scarcity of the fruiting bodies [7], and general appreciation is bound to be af- fected by availability and ease of access to the resource. When some taxa are rare or only available in more re- mote areas, people may find it difficult to go far to col- lect them. The result is that mushroom diets can and will shift as a mere result of access and distance to suit- able sites [7]. We have no data from eventual hunter-gatherers in the region, but the main reason why people from the Tshopo show a preference for saprotrophic fungi is that these species are found close to the villages and nearby recently cut forests. Because of frequent rains and avail- ability of dead wood, saprotrophic taxa also manage to produce fruit bodies all year round. In contrast, EcM fungi occur abundantly in remote and pristine (or nearly pristine) forests with older growth of Gilbertiodendron dewevrei, Brachystegia laurentii,orUapaca spp. [19]. With the exception of Cantharellus rufopunctatus, pro- Fig. 3 Number of ectomycorrhizal and saprotrophic fungi within ducing fruit bodies at least from April to December [9], monodominant and mixed forests. NS no significant difference; most of the edible EcM taxa can only be found during a ***highly significant difference short period of time. Milenge Kamalebo and De Kesel Journal of Ethnobiology and Ethnomedicine (2020) 16:8 Page 9 of 13

Table 2 Edible fungi local names and list of fungi eaten by informants from each ethnic group Species Local names Ethnic groups Kumu Lokele Ngando Ngelema Topoke Turumbu Agaricus bambusicola Heinem. NA − ++ + + + Agaricus crocopeplus Berk. & Broome NA − ++ + + + Amanita annulatovaginata Beeli NA − ++ + + + Amanita echinulata Beeli NA + + + + + + Amanita pudica (Beeli) Walleyn NA + + + + + + Amanita robusta Beeli Balenda1 ++ + + + + Armillaria heimii Pegler Baselekoko3 ++ + + + + Auricularia cornea Ehrenb. Batele1,2,3,4,5,6 ++ + + + + Auricularia delicata (Mont.ex Fr.) Henn. Batele1,2,3,4,5,6 ++ + + + + Calyptella longipes (Cooke & Massee) W.B. Cooke Bape-bape6 − ++ − ++ Cantharellus congolensis Beeli Bolingobopii6 ++ + + + + Cantharellus conspicuus Eyssartier, Buyck & Verbeken NA −− − − − − Cantharellus densifolius Heinem. NA + + + + + + Cantharellus floridulus Heinem. Solosolo3 ++ + + + + Cantharellus incarnatus (Beeli) Heinem. NA −− − − − − Cantharellus longisporus Heinem. Baolo-bandei6, Abamongo1 ++ + + + + Cantharellus luteopunctatus (Beeli) Heinem. Baolo-bandei6, Abamongo1 ++ + + + + Cantharellus miniatescens Heinem. Lilenda-linenu6, Agbalo-gbalo4 ++ + + + + Cantharellus aff. ruber Heinem. NA + + + + + + Cantharellus rufopunctatus (Beeli) Heinem. Bengole-bonyele6 ++ + + + + Cookeina speciosa (Fr.) Dennis Batoyi bangwa’a2 −− + −−+ Cookeina tricholoma (Mont.) Kuntze Batoyi bangwa’a2 −− − − − − Coprinellus disseminatus (Pers.) J.E. Lange Baku’kulekule6 − ++ + + + Coprinus plicatilis (Curtis) Fr. NA −− − − − − Cotylidia aurantiaca (Pat.) A. L. Welden NA −− + −−+ Dacryopinax spathularia (Schwein.) G.W. Martin NA −− − − − − Favolus tenuiculus (P. Beauv.) Fr. NA −− + −−+ Gerronema hungo (Henn.) Degreef & Eyi NA −− − − − − Gymnopilus zenkeri (Henn.) Singer Bokoo’kina6 ++ + + + + Hygrocybe cantharellus (Schwein.) Murrill Adanopila4 − ++ + + + Hygrocybe coccinea (Schaeff.) P. Kummer NA −− − − − − Hymenagaricus luteolosporus Heinem.& Little Flower Baolo ba’ongolo6, Tongwengwe4 − ++ + + + Hypholoma subviride Berk. & M.A. Curtis NA −− − − − − Lactarius acutus Heim Mongo5 − ++ + + + Lactifluus annulatoangustifolius (Beeli) Buyck Belaa’tala3 − ++ + + + Lactifluus gymnocarpus (R. Heim ex Singer) Verbeken NA −− − − − − Lactifluus heimii (Verbeken) Verbeken NA −− − − − − Lactifluus pelliculatus (Beeli) Buyck Batina lya bakuu6 −− − − − − Lentinus squarrosulus Mont. Bengole6 ++ + + + + Lepista rhodotoides Singer Bongolototo6, Baselesele2 − ++ + + + Lepista sordida (Schumach.) Singer NA −− − − − − Leucocoprinus cepaestipes (Sowerby) Pat Mbumbuke6 − ++ − ++ Leucocoprinus discoideus (Beeli) Heinem NA −− − − − − Milenge Kamalebo and De Kesel Journal of Ethnobiology and Ethnomedicine (2020) 16:8 Page 10 of 13

Table 2 Edible fungi local names and list of fungi eaten by informants from each ethnic group (Continued) Species Local names Ethnic groups Kumu Lokele Ngando Ngelema Topoke Turumbu Macrolepiota dolichaula (Berk. & Broome) Pegler & R.W. NA −− − − − − Rayner Marasmiellus inoderma (Berk.) Singer NA −− − − − − Marasmius arborescens (Henn.) Beeli Twendenda5, Seeliko3 ++ + + + + Marasmius bekolacongoli Beeli Bakoko wa kombe6, Toko kombe2 ++ + + + + Marasmius buzungolo Singer Mongala6, Ndundo3,5 ++ + + + + Marasmius confertus Berk. & Broome Mongala6 ++ + + + + Neonothopanus hygrophanus (Mont.) De Kesel & NA −− − − − − Degreef Paxillus brunneotomentosus Heinem. & Rammeloo NA −− − − − − Pleurotus cystidiosus O.K. Mill. Bengole6,2,5 ++ + + + + Pleurotus flabellatus (Berk. & Br.) Sacc. Bengole6,2,5 ++ + + + + Pleurotus tuber-regium (Fr.) Fr. Bengole ya makasi6 ++ + + + + Polyporus arcularius (Batsch) Fr. NA −− − − − − Psathyrella candolleana (Fr.) Maire NA −− − − − − Rubinoboletus luteopurpureus (Beeli) Heinem. & Lofyongi lonyele6 − ++ + + + Rammeloo Russula annulata R. Heim Lilianga6 − ++ + + + Russula inflata Buyck Balenda ya moindo6 − ++ − ++ Russula meleagris Buyck Nsalanka’ngaa6 ++ + + + + Russula porphyrocephala Buyck Nsololo6 ++ + + + + Russula pruinata Buyck Balenda ya kulokoko6 − ++ + + + Russula roseostriata Buyck NA −− − − − − Russula sese Beeli NA −− − − − − Russula sesemoindu Beeli Malebadja6 − ++ + + + Russula striatoviridis Buyck NA − ++ − ++ Schizophyllum commune Fr. Bokotoko6, Tukumu2, Bokokola1,3,4,5 ++ + + + + Tapinella panuoides (Fr.) E.-J. Gilbert Tukutuno6 − ++ − ++ Termitomyces robustus (Beeli) Heim Limusula6, Libusula5, Bululu3, ++ + + + + Nku’suusu1 Termitomyces singidensis Saarim. & Härk. Limusula6, Libusula5, Bululu3 − ++ − ++ Termitomyces striatus (Beeli) Heim Limusula6, Libusula5, Bululu3 ++ + + + + Tremella fuciformis Berkeley NA −− − − − − Tricholomopsis aurea (Beeli) Desjardin & B.A. Perry. NA + + + + + + Trogia infundibuliformis Berk. & Br. NA + + + + + + Tylopilus balloui (Peck) Singer NA −− − − − − Volvariella parvispora Heinem. NA −− − − − − Volvopluteus gloiocephalus (DC) Vizzini NA −− − − − − Xerocomus spinulosus Heinem. & Gooss.-Font. Kwekwele6 −− + −−+ 1Kumu, 2Lokele, 3Ngando, 4Ngelema, 5Topoke, 6Turumbu (mother tongues); “+” indicates edible fungi and “−” indicates inedible fungi by local ethnic groups’ informants; NA not applicable

Conclusion Although not formally measured, there is little doubt that The results from this study reveal that rainforests from the the local populations collect and use only a small fraction of Yangambi Biosphere reserve and Yoko forest reserve of the the naturally produced quantity of edible fungi. Most of the Tshopo provide a substantial number of edible fungi. locally used taxa are saprotrophic and easily accessed from Milenge Kamalebo and De Kesel Journal of Ethnobiology and Ethnomedicine (2020) 16:8 Page 11 of 13

Table 3 Mean numbers of edible fungi eaten by informants from each ethnic group (tribe) ± standard deviation (SD) Ethnic Fungal trophic groups Kruskal-Wallis groups Ectomycorrhizal Saprotrophic Termitophilic p value Significance fungi, mean ± fungi, fungi, SD mean ± SD mean ± SD Turumbu 8 ± 3 13 ± 3 2 ± 1 < 0.001 *** Lokele 7 ± 3 12 ± 3 2 ± 1 < 0.001 *** Topoke 7 ± 3 13 ± 3 2 ± 1 < 0.001 *** Ngando 19 ± 3 24 ± 4 2 ± 1 < 0.001 *** Ngelema 10 ± 3 12 ± 2 2 ± 1 < 0.001 *** Kumu 8 ± 2 11 ± 2 2 ± 1 < 0.001 *** p value < 0.001 < 0.001 > 0.05 Significance *** *** NS The two last columns and lines indicate the p value and significance level of the Kruskal-Wallis test, respectively (NS no significant difference, ***very high significant difference) either litter or dead wood. It is the availability, abundance, Some locally considered inedible EcM fungal taxa are and proximity of this resource that makes saprotrophic fungi in fact perfectly edible and used for food elsewhere in from anthropized environments (secondary forests, clearings, Africa. More research needs to be done, but assessing fallow, farmland) most attractive and preferred. As EcM the natural productivity of edible fungi from the rain- fungi are harder to obtain and less frequent, they are a poorly forests should help us better understand what these known and underutilized food source. Whereas in the Zam- fungi can mean as an ecosystem service to local liveli- bezian region, the major local interest for edible EcM fungi hoods. In combination with the measured interest of can be used as a strong incentive for forest conservation; our local people for this resource, one should be able to de- observations indicate that this is not the case in the Tshopo velop systems to promote the use of this largely un- region. tapped resource.

Fig. 4 Four of the most important edible saprotrophic fungi in Tshopo province. a Auricularia delicata. b Lentinus squarrosulus. c Marasmius buzungolo. d Schizophyllum commune Milenge Kamalebo and De Kesel Journal of Ethnobiology and Ethnomedicine (2020) 16:8 Page 12 of 13

Abbreviations 5. Degreef J, Malaisse F, Rammeloo J, Baudart E. Edible mushrooms of the DR Congo: Democratic Republic of the Congo; EcM: Ectomycorrhizal (fungi, Zambezian woodland area. Nutritional and ecological approach. Biotechnol forests, trees); EMCS: Edible mushrooms’ cultural significance; NTFP: Non- Agron Soc Environ. 1997;1(3):221–31. timber forest product; Sapr: Saprotrophic fungi; Term: Termitophilic fungi 6. Khatun S, Islam A, Cakilcioglu U, Guler P, Chatterjee NC. Nutritional qualities and antioxidant activity of three edible oyster mushrooms (Pleurotus spp.). – Acknowledgements NJAS- Wageningen J Life Sci. 2015;72(73):1 5. The first author acknowledges the PhD scholarship granted by the Center for 7. Degreef J, Demuynck L, Mukandera A, Nyirandayambaje G, Nzigidahera B, International Forestry Research (CIFOR) at the University of Kisangani (DR De Kesel A. Wild edible mushrooms, a valuable resource for food security Congo) and the research grant from the International Foundation for and rural development in Burundi and Rwanda. Biotechnol Agron Soc – Science (IFS/ I-3-D-6072-1) that supported respectively fungal sampling ex- Environ. 2016;20(4):441 52. ’ pedition and ethnomycological data collection. Many thanks to the Belgian 8. Buyck B. Flore illustrée des champignons d Afrique centrale: Russula II – Global Taxonomy Initiative of the CEBioS program for supporting the taxo- (). Bull Jard Bot Nat Belg. 1994a;16:411 542. nomic analysis for species identification at the Meise Botanic Garden in 9. De Kesel A, Amalfi M, Kasongo B, Yorou NS, Raspé O, Degreef J, Buyck B. Belgium. A. De Kesel acknowledges the Belgian Federal Science Policy Office New and interesting Cantharellus from tropical Africa. Cryptogam Mycol. (BELSPO) for financing his fieldwork in Yangambi through the project 2016;37:283–327. https://doi.org/10.7872/crym/v37.iss3.2016.283. COBIMFO (Congo basin integrated monitoring for forest carbon mitigation 10. De Kesel A, Malaisse F. Edible wild food: fungi. In: Malaisse F, editor. How to and biodiversity). Ratos RISASI, Jules BOMBILE, Antoine MOTOSIA, and Michel live and survive in Zambezian open forest (Miombo ecoregion). Gembloux: MBASI are acknowledged for the assistance and guidance during the sam- Presses Agronomiques de Gembloux; 2010. p. 41–56. pling expedition and ethnomycological survey campaign. 11. Härkönen M, Niemelä T, Mbindo K, Kotiranta H, Piearce G. Zambian mushrooms and mycology. Norrlinia 29; 2015. 12. Härkönen M, Niemelä T, Mwasumbi L. Tanzanian mushrooms - edible, Authors’ contributions harmful and other fungi. Norrlinia 10; 2003. HMK designed the study; supplied the objectives; collected the data; did the 13. Malaisse F. Se nourrir en forêt claire Africaine. Approche écologique et species identification, data analysis, and interpretation; and wrote the first nutritionnelle. Faculté des Sciences agronomiques de Gembloux; 1997. manuscript draft. ADK supplied the tools for identification; did the data collection, species identification (macro- and microscopy), and laboratory 14. Malaisse F, Guy P. Edible wild vegetable products in the Zambezian woodland area: a nutritional and ecological approach. Ecol Food Nutr. 1985; work; and is the co-writer and responsible for the corrections to the English – text. Both authors read and approved the final manuscript. 18:43 82. 15. Boni S, Yorou NS. Diversité et variabilité inter-ethnique dans la consommation de champignons sauvages de la région de N’Dali au Bénin. Funding Tropicultura. 2015;33(4):266–76. Center for International Forestry Research (CIFOR), Belgian Federal Science 16. De Kesel A, Codjia JTC, Yorou NS. Guide des champignons comestibles du Policy Office (BELSPO), and International Foundation for Science (IFS) Bénin. COCO MULTIMEDIA; 2002. 17. Yorou SN, De Kesel A. Connaissances ethnomycologiques des peuples Availability of data and materials Nagot du centre du Bénin (Afrique de l’Ouest). In: Robbrecht E, Degreef J, The data that support the findings of this study have been deposited at the Friis I, editors. Plant systematics and phytogeography for the understanding herbarium of the Meise Botanic Garden in Belgium (BR) through the of African biodiversity, Proceedings of the XVIth AETFAT Congress 2000 reference codes ADK (for the collection made by André De Kesel) and MKH (Meise, National Botanic Garden of Belgium), vol. 71; 2001. p. 627–37. Syst. (for the collection made by Milenge Kamalebo Héritier). Some data are Geogr. Pl. accessible on https://www.efta-online.org/. The overall data are available 18. Eyi-Ndong H, Degreef J, De Kesel A. Champignons comestibles des forêts denses from the authors. d’Afrique centrale. Taxonomie et identification. Abc Taxa volume 10; 2011. 19. Milenge KH, Nshimba SMH, Masumbuko NC, Nabahungu NL, Degreef J, De Ethics approval and consent to participate Kesel A. Host plants and edaphic factors influence the distribution and Not applicable diversity of ectomycorrhizal fungal fruiting bodies within rainforests Tshopo, Democratic Republic of the Congo. Afr J Ecol. 2019;00:1–13. https://doi.org/ 10.1111/aje.12595. Consent for publication 20. Lejoly J, Ndjele M-B, Geerinck D. Catalogue-flore des plantes vasculaires des Not applicable districts de Kisangani et de la Tshopo (R D Congo). Taxonomania. 2010; 4(30):1–308. Competing interests 21. Amani IC. Vegetation patterns and role of edaphic heterogeneity on plant The authors declare that they have no competing interests. communities in semi-deciduous forests from the Congo basin. PhD thesis, Université Libre de Bruxelles; 2011. p. 198. 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